Effects of feeding different amounts of supplemental glycerol on ruminal environment and

digestibility of lactating dairy cows

Principio del formulario

J. Boyd

Affiliations

Department of Animal and Dairy Science, The University of Georgia, Tifton 31793

US Dairy Forage Research Center, 1925 Linden Drive, West Madison, WI 53706.

Corresponding author:

J.K. Bernard

Affiliations

Department of Animal and Dairy Science, The University of Georgia, Tifton 31793

J.W. West

Affiliations

Department of Animal and Dairy Science, The University of Georgia, Tifton 31793

Final del formulario

Received 22 May 2012; accepted 2 October 2012. published online 26 November 2012.

Abstract

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Article Outline

Abstract
Introduction

Materials and Methods

Cows and Management

Experimental Treatments and Design

Data Collection

Digestibility Study

Statistical Analysis

Results and Discussion

Conclusions

Acknowledgments

References

Copyright

Abstract

A replicated 3 × 3 Latin square study was conducted to evaluate the effects of replacing a portion
of ground corn in the diet with dietary glycerol on rumen environment, blood metabolites, and
nutrient digestibility. Six rumen-cannulated Holstein cows, averaging 56 ± 18 DIM and 38.0 ± 8.2
kg of milk/d, were used in the study. Experimental periods included 3 wk for treatment
adjustment period followed by 1 wk for data collection. Diets were corn silage based and balanced
to be isocaloric and isonitrogenous. Treatments were 0 g of glycerol/d (control, CON), 200 g of
glycerol/d (G2), and 400 g of glycerol/d (G4). Dry matter intake (DMI) decreased as the amount of
glycerol fed increased. Milk yield was higher for CON and G2 cows in comparison with G4 cows.
Milk fat percentage and yield were reduced when glycerol was fed compared with CON cows but
increased the milk protein percentage at the highest concentration of dietary glycerol. These
changes resulted in decreased energy-corrected milk yield and efficiency (milk/DMI) in diets
supplemented with G4 compared with CON. No differences were observed in ruminal pH and
ammonia concentrations. Molar proportions of acetate, valerate, and acetate:propionate ratio
decreased, whereas propionate increased as the amount glycerol fed increased. Molar proportions
of butyrate were greatest when glycerol was included in the diet compared with CON. Nutrient
intake and digestion were not different among treatments. Results of this trial suggest that
feeding increasing amounts of glycerol may decrease DMI and alters ruminal fermentation,
resulting in reduced yield of milk, fat, and energy-corrected milk.

Key words: dietary glycerol, volatile fatty acid, efficiency

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Introduction

The growth of the ethanol industry has increased the need for alternatives to corn in lactating cow
diets. Biodiesel production has increased in the United Sates from 690 million gallons in 2009 to
802 million gallons in 2011 (Biofuels Digest, 2011). Industry growth is expected to increase
availability and promote favorable pricing of glycerol, which is a byproduct of base-catalyzed
transesterification of oil in the formation of methyl and ethyl FA esters in the production of
biodiesel (Thompson and He, 2006) and the main byproduct of ethanol fermentation processing
(Michnick et al., 1997). Purified glycerol is used in the food, pharmaceutical, and cosmetic
industries, but the cost of refining crude glycerol to a high purity is cost prohibitive for many small
refining operations (Pachauri and He, 2006). This leads to potentially a large source of crude
glycerol available to livestock producers. According to the Food and Drug Administration (FDA,
2006), glycerol is recognized as a safe feed ingredient in animal diets. Glycerol is an odorless,
colorless, hygroscopic, sweet-tasting liquid that has the potential to replace corn in the diet and
has been predicted to have a feed value of 100 to 120% of corn, making it a viable alternative to
corn (Hippen et al., 2008). Linke et al. (2004) calculated the NEL value of glycerol to be
approximately 1.05 Mcal/kg, which is 20% greater than corn. Schröder and Südekum (1999)
calculated the energy density of glycerol to range between 0.90 to 1.03 Mcal of NEL/kg. Also,
several studies have reported that glycerol enhances rumen fermentation and improves feed
efficiency (Garton et al., 1961; Dirksen et al., 1985; Hippen et al., 2008). Dietary glycerol seems to
be extensively fermented to propionate by ruminal bacteria (Czerkawski and Breckenridge, 1972;
Remond et al., 1993; Bergner et al., 1995), although level of feeding and method of delivery may
affect the amount of glycerol that escapes fermentation (Kijora et al., 1998; DeFrain et al., 2004).

The increased availability may create an opportunity to use glycerol as an energy source in dairy
rations and has led to a corresponding decline in price projections (Yazdani and Gonzalez, 2007).
These changes may favor the use of glycerol in rations fed to livestock, especially with the rising
corn costs and more crops being diverted to biofuel production.

Previously, glycerol has been a cost-prohibitive addition to the diet because of its value in other
fields, but with growing biodiesel production and corn costs, the addition of glycerol to the ration
is a possible choice for many producers. The objective of this study was to determine the effects of
feeding diets with different levels of glycerol on rumen function, blood metabolites, and nutrient
digestibility in lactating Holstein cows.

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Materials and Methods

The study was conducted May to July of 2008 at the University of Georgia, Tifton Campus Dairy
Research Center. The protocol for the trial was approved by the University of Georgia Institutional
Animal Care and Use Committee.
Cows and Management

Six early-lactation rumen-cannulated Holstein cows (4 multiparous and 2 primiparous) were used
in the replicated 3 × 3 Latin square design study. Cows averaged 56 ± 18 DIM, 38.0 ± 8.2 kg/d of
milk, 3.77 ± 0.97% milk fat, and 2.72 ± 0.2% milk protein at the beginning of the study. Cows were
housed in a freestall barn with access to individual freestalls, free access to water, and fed behind
Calan doors (American Calan Inc., Northwood, NH). Training for Calan doors was initiated in early
April, and the study began in early May and continued until the end of July. Cows were cooled
using fans and high-pressure misters. All diets contained 15 g/907 kg of DM of Rumensin (Elanco
Animal Health, Greenfield, IN).

Feed was mixed and delivered once daily. The amount of feed offered was adjusted each day to
achieve 7 to 10% refusals. Diets were balanced to be isocaloric and isonitrogenous (Table 1). Cows
were milked twice daily at 0400 and 1500 h during the study.

Table 1. Ingredient and chemical composition of experimental diets containing supplemental

glycerol1

Item Diet2

CON G2 G4

Ingredient, % of DM

Corn silage 38.5 38.5 38.5

Alfalfa hay 10.6 10.6 10.6

Whole cottonseed 1.82 1.82 1.82

Ground corn 23.9 22.9 22.1

Wet brewers grain 12.1 12.1 12.1

Concentrate mix3 13.2 13.2 13.2

Glycerol 0 0.78 1.57

Item Diet2

CON G2 G4

DM,4 % 50.0 ± 0.5 49.3 ± 0.8 49.2 ± 0.6

Chemical composition,4 % of DM

NDF 39.6 ± 1.8 40.9 ± 1.9 39.7 ± 1.5

ADF 26.1 ± 1.1 27.1 ± 1.6 26.3 ± 1.1

CP 17.2 ± 0.5 16.6 ± 0.3 16.7 ± 0.6

RUP5 7.14 6.86 6.89

RDP5 10.05 9.74 9.81

Ether extract 4.4 ± 0.3 4.7 ± 0.3 4.7 ± 0.6

1Samples were collected 4 times during the collection week and compiled for analysis (n = 3).

2CON = control: no glycerol added; G2 = control + 200 g of supplemental glycerol/d; G4 = control

+ 400 g of supplemental glycerol/d.

3Composition of concentrate mix (% of DM): 2.04% Pro-Lak (H. J. Baker and Brother Inc.,
Westport, CT), 6.06% soybean meal 48, 0.151% Ca 17%:P 21%, 0.182% potassium magnesium
sulfate, 0.454% potassium bicarbonate, 0.151% limestone, 0.151% urea 45%N, 0.027% Availa-4
(Zinpro Corp., Eden Prairie, MN), 0.154% MgO, 0.227% salt; 0.530% Na bicarbonate, 0.188% yeast,
0.121% trace mineral premix (2.91% Ca; 0.38% Mg; 1.76% S; 144 mg/kg of Co; 9,523 mg/kg of Cu;
1,465 mg/kg of Fe; 842 mg/kg of I; 28,617 mg/kg of Mn; 220 mg/kg of Se; 25,343 mg/kg of Zn),
0.01% vitamin premix (4,210,830 IU/kg of vitamin A; 1,684,330 IU/kg of vitamin D; 21,045 IU/kg of
vitamin E), and 0.229% Rumensin 3 (Elanco Animal Health, Greenfield, IN).

4Results are presented as means ± SD.

Item Diet2

CON G2 G4

5Calculated using the Spartan ration program (Michigan State University, East Lansing).

Cows were ranked by average daily milk yield, stage of lactation, and ECM during the
standardization period, blocked into pairs by rank, and each pair was assigned randomly to 1 of
the 3 experimental diets within block and progressed through the three 4-wk periods until
exposed to all treatments.

Experimental Treatments and Design

The experimental design for this study was a replicated 3 × 3 Latin square. Each period consisted
of 3 wk for treatment adjustment followed by 1 wk for data collection. Dietary treatments
consisted of a control without supplemental glycerol (CON), control plus 200 g of supplemental
glycerol/d (G2), and control plus 400 g of supplemental glycerol/d (G4; Table 1). Glycerol was
mixed into the TMR using a top-loading Super Data Ranger (American Calan Inc.) with whole
cottonseed as a carrier. The glycerol was added to the whole cottonseed and mixed and the
coated cottonseed was then added to the TMR in place of ground corn. Glycerol contained 80 to
85% glycerol with no more than 14% moisture, 7% sodium chloride, and methanol content of 18
ppm (analysis provided by ADM Nutrition Alliance Inc., Quincy, IL).

Data Collection

The amount of feed offered and refused was recorded daily during the 7-d collection week. Milk
yield was recorded at each milking (2×) using electronic meters (Alpro; DeLaval, Kansas City, MO)
and summed daily during the collection weeks. Milk samples were collected from 4 consecutive
milkings each collection period and analyzed for concentrations of fat, protein, and SCC (Southeast
Milk Inc., Belleview, FL). Energy-corrected milk yield was calculated as ECM = (0.327 × kg of milk) +
(12.95 × kg of fat) + (7.65 × kg of protein) (Tyrrell and Reid, 1965).

Blood samples from the coccygeal vessel were collected once during the each collection week for
analysis of serum glucose and urea N using a Boehringer Mannheim/Hitachi 912 automated
chemistry analyzer (Roche Laboratory Systems, Indianapolis, IN). Rumen samples were collected
on d 3 of each collection week at 0, 2, 4, 6, 8, and 10 h after feeding. Approximately 50 mL of
ruminal fluid was collected and strained through 3 layers of cheesecloth and immediately analyzed
for pH and ammonia levels. A 10-mL subsample was immediately mixed with 2 mL of
metaphosphoric acid (25% wt/vol), centrifuged at 10,000 × g for 10 min, and the supernatant was
collected and frozen for later analyses of VFA (Erwin et al., 1961) using a Thermo Scientific Trace
GC Ultra (Thermo Fisher Scientific Inc., Bellefonte, PA).
Samples of TMR, refusals, and ingredients were collected 4 times during the collection week. The
DM content was determined by drying in a forced-air oven at 55°C for 48 h. Samples were
composited by period and ground to pass through a 1-mm screen using a Wiley mill (Thomas
Scientific, Swedesboro, NJ) for analysis of DM, CP, ether extract (AOAC, 1990), NDF (Van Soest et
al., 1991), and ADF (AOAC, 1990).

Digestibility Study

Cows were fed 23 g/d (DM basis) of Cr2O3 mixed into the experimental diet beginning 10 d before
the 4-d collection period each collection week. Powdered Cr2O3 was mixed with ground corn as a
carrier at a rate of 3.82 g of corn/kg of DM in an upright mixer. The Cr2O3 corn blend was added
to the diet and mixed daily. Daily feed and ort samples were collected during the 4-d collection
period. Fecal grab samples were collected during the collection period at 12-h intervals, with the
collection time advancing by 3 h each day to represent 0300, 0600, 0900, 1200, 1500, 1800, 2100,
and 2400 h. Fecal samples were frozen at −5°C until composited by cow. Samples of feed, orts, and
feces were compiled by cow and dried at 55°C and ground to pass through a 1-mm screen using a
Wiley mill and stored for analysis of CP, NDF, and ADF as described previously. Chromium content
of feed, orts, and feces was determined using atomic absorption spectrophotometry (AAnalyst
100/300; Perkin Elmer, Bellefontaine, PA) following wet ashing (Ferret et al., 1999).

Statistical Analysis

Data were analyzed as a 3 × 3 Latin square design using PROC MIXED of SAS version 9.1 (SAS
Institute, 2004). Significance was defined as P < 0.05 and trends as 0.05 ≤ P ≤ 0.15. Values are
reported are least squares means and associated standard errors. Model 1 was used for analysis of
intake, production, blood, and apparent digestibility. Model 2 was used for analysis of VFA, pH,
and ammonia and included effects of time, period, and treatment. Interactions tested were hour
by treatment for ruminal data. Sample time was included as a repeated measure for model 2.
When significance was determined, pairwise t-tests with a Tukey adjustment were performed
between treatment means using the PDIFF statement in SAS. The models were as follows:

where Yijmk and Yijln = variable responses, μ = general mean, βi = fixed effect of treatment (i = 1,
. . . , 3), Pj = fixed effect of period (j = 1, . . . , 3), Ck = fixed effect of cow (k = 1, . . . , 6), Hl = fixed
effect of hour (l = 0, . . . , 10), βHil = interaction of treatment and hour, and εijmk and εijkln =
residual errors.

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Results and Discussion

The chemical composition of the experimental diets is reported in Table 1. Nutrient concentrations
were similar for all diets except that concentrations of ether extract were slightly higher when
glycerol was included in the diet.

Milk yield was reduced (P = 0.004) by 1.8 and 2.4 kg/d for G4 compared with CON and G2.
Inclusion of glycerol in the diet resulted in a reduction in milk fat percentage (P = 0.004) and yield
(P = 0.0009) compared with CON. In contrast, milk protein percentage increased (P = 0.0001) at
the higher level of glycerol supplementation compared with CON and G2. These results are in
contrast to earlier work reported by Donkin and Doane (2007) in which no differences were
observed in milk yield or composition when glycerol replaced corn in the diet. Chung et al. (2007)
included dried glycerol (food grade, 65% glycerol) in the diet of transition cows and also reported
no effect on milk yield or composition. Linke et al. (2004) fed glycerol at the rate of 498 or 998 g/d
in the diet of mid-lactation Holstein and Brown Swiss cows in a 3 × 3 Latin square, with no effect
on DMI, milk yield, or FCM, but observed an improvement in feed efficiency with the inclusion of
glycerol with milk to feed ratios of 1.46, 1.59, and 1.60 for the control, 498 and 998 g/d,
respectively. Feed efficiency (milk/DMI) was reduced (P = 0.05) for G4 compared with G2 and
CON. Also, ECM was reduced (P = 0.003) for G4 compared with CON and a tendency (P = 0.10) for
lower efficiency (ECM/DMI) was found in the current study. These results are in agreement with
DeFrain et al. (2004) who reported reduced ECM when glycerol was fed. No differences were
observed among treatments in concentrations of serum glucose or urea N, although a trend (P =
0.09) for reduced serum urea N was observed with increasing amounts of glycerol in the diet
(Table 2).

Table 2. Performance and blood metabolites of lactating Holstein cows fed diets supplemented
with different amounts of supplemental glycerol (n = 72 milk samples; n = 18 serum samples)

Item Diet1 SE P-value

CON G2 G4

Milk, kg/d 37.9a 37.3a 35.5b 0.52 0.004

Fat, % 3.46a 3.31b 3.35b 0.03 0.004

Fat, kg/d 1.33a 1.24b 1.19b 0.03 0.0009

Protein, % 2.76a 2.75a 2.81b 0.005 0.0001

Protein, kg/d 1.04 1.02 1.00 0.02 0.06

Item Diet1 SE P-value

CON G2 G4

ECM, kg/d 37.6a 36.1ab 34.6b 0.61 0.003

Efficiency

ECM/DMI 1.54 1.55 1.48 0.02 0.10

Milk/DMI 1.55ac 1.60ab 1.51c 0.03 0.05

Serum metabolite

Glucose, mg/dL 62.7 62.5 63.3 1.3 0.89

Urea N, mg/dL 21.7 21.8 20.5 0.4 0.09

a–cMeans within a row with unlike superscripts differ (P < 0.05).

1CON = control: no glycerol added; G2 = control + 200 g of supplemental glycerol/d; G4 = control

+ 400 g of supplemental glycerol/d.

No differences were observed for the interaction of treatment and sampling time; thus, only the
main effects are reported. No differences were observed among treatments for ruminal pH
(average 6.1), but a weak trend (P = 0.14) existed for decreased ruminal ammonia concentrations
with the addition of glycerol to the diet. Ruminal ammonia concentrations were 12.1, 11.7, and 9.8
for CON, G2, and G4, respectively (Table 3). No differences were observed in total VFA
concentration among treatments. Molar proportions of acetate, valerate, and acetate:propionate
ratio decreased (P = 0.0001), whereas molar proportions of propionate (P = 0.0002) increased as
the amount of glycerol fed increased. Molar proportions of butyrate were highest (P = 0.0001) for
diets supplemented with glycerol compared with CON. No differences were observed in molar
proportions of isobutyrate or isovalerate.

Table 3. Ruminal pH and VFA concentrations of lactating Holstein cows fed diets supplemented
with different amounts of supplemental glycerol (n = 108 rumen fluid samples)1
Item Diet2 SE P-value

CON G2 G4

pH 6.1 6.0 6.1 0.04 0.64

Ammonia, % 12.1 11.7 9.8 0.86 0.14

Total VFA, mmol 46.7 52.2 49.0 2.3 0.24

Composition, %

Acetate 61.1a 59.1b 57.6c 0.39 0.0001

Propionate 23.3a 24.1b 25.4c 0.35 0.0002

Butyrate 11.4a 12.3b 12.4b 0.15 0.0001

Isobutyrate 1.3 1.2 1.2 0.06 0.91

Isovalerate 1.5 1.5 1.5 0.04 0.53

Valerate 1.5a 1.7b 1.8b 0.05 0.0001

Acetate:propionate 2.7a 2.5b 2.3c 0.05 0.0001

a–cMeans within a row with unlike superscripts differ (P < 0.05).

1No treatment by sampling time interactions (P > 0.10).

2CON = control: no glycerol added; G2 = control + 200 g of supplemental glycerol/d; G4 = control

+ 400 g of supplemental glycerol/d.

The effects of supplemental glycerol on ruminal VFA concentrations are in agreement with
previous reports. Schröder and Südekum (1999) examined the effects of feeding glycerol on
ruminal fermentation using ruminally cannulated steers and reported that although the addition
of glycerol did not affect diet digestibility, it did decrease the acetate:propionate ratio, linearly
increased ruminal butyrate concentrations, and stimulated increased water intake. Schröder and
Südekum (1999) concluded that the inclusion of dietary glycerol would be beneficial to the dairy
cow because it would increase ruminal propionate, increasing the supply of gluconeogenic
substrate to the liver. Also, the increased ruminal butyrate would support the growth of ruminal
epithelial tissue and possibly increase nutrient absorption from the rumen as proposed by Dirksen
et al. (1985).

DeFrain et al. (2004) conducted a transition cow study with the inclusion of glycerol. Those
researchers reported that rumen fluid collected postpartum from cows offered glycerol had a
greater total VFA concentration, increased molar proportions of propionate, and decreased
acetate:propionate ratio compared with the control. Butyrate concentrations also showed a
tendency to increase linearly in cows fed glycerol treatment compared with CON.

Linke et al. (2004) reported an increase in molar proportions of propionate and butyrate with
increasing glycerol without any difference in DMI, milk yield, or FCM. They also reported molar
proportions of acetate decreased when cows were provided supplemental glycerol through
drenching or mixing in their diet. Molar proportions of propionate and butyrate increased with
either route of administration with peak concentration observed 4 h after consumption.
Concentrations of glycerol in blood plasma increased when cows were drenched or tubed but not
when they were fed glycerol. Plasma insulin and BHBA concentrations increased when cows were
drenched or tubed compared with those fed glycerol in their diets. These researchers concluded
that for glycerol to be gluconeogenic it must be delivered in water to associate with the liquid
fraction of the rumen contents or be able to bypass the rumen and be absorbed as glycerol and
converted to glucose by the liver. Glycerol that is available to the rumen microbes is highly
fermentable and is converted to propionic and butyric acids. Garton et al. (1961) conducted in
vitro incubations of glycerol and observed that almost 25% of glycerol had disappeared after 2 h
and almost 90% of the glycerol was undetectable after an 8-h incubation.

The DMI was reduced (P = 0.003) with the inclusion of glycerol for both G2 and G4 compared with
CON (Table 4). This is in contrast to a previous trial (Boyd et al., 2011) in which no differences in
DMI were observed when lactating cows were fed diets supplemented with up to 400 g of
glycerol/d for 10 wk. Other researchers (DeFrain et al., 2004; Linke et al., 2004) have not observed
any effect on DMI with the inclusion of glycerol in the diet. However, no differences in nutrient
intake or apparent digestibility (Table 4) were observed among treatments. These results are in
agreement with previous research reported by Schröder and Südekum (1999) and Donkin and
Doane (2007) who reported no effects on nutrient digestibility with the inclusion of glycerol in the
diet.

Table 4. Nutrient intake and apparent digestibility of lactating Holstein cows fed diets
supplemented with different amounts of supplemental glycerol (n = 108 intake observations; n =
144 fecal samples)
Item Diet1 SE P-value

CON G2 G4

Intake, kg/d

DM 24.3a 23.1b 23.4b 0.24 0.003

CP 3.5 3.1 3.1 0.1 0.21

NDF 7.8 7.4 7.5 0.4 0.70

ADF 5.2 4.9 4.8 0.3 0.60

Apparent digestibility, %

DM 68.5 66.9 65.6 2.3 0.66

CP 71.8 69.1 68.2 1.8 0.36

NDF 47.5 46.3 48.6 3.5 0.90

ADF 44.9 43.6 40.9 4.1 0.78

a,bMeans within a row with unlike superscripts differ (P < 0.05).

1CON = control: no glycerol added; G2 = control + 200 g of supplemental glycerol/d; G4 = control

+ 400 g of supplemental glycerol/d.

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Conclusions

Inclusion of dietary glycerol in the ration of dairy cows affected milk yield and components, ECM,
and ruminal fermentation patterns. The effects on ruminal fermentation are in agreement with
several earlier studies. Addition of rumen-fermentable glycerol decreased molar proportions of
acetate and acetate:propionate ratio and increased molar proportions of propionate and butyrate,
which should increase the supply of gluconeogenic precursors and support the growth of ruminal
epithelial tissue and potentially increase nutrient absorption from the rumen.

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Acknowledgments

The authors thank ADM Alliance Nutrition Inc. (Quincy, IL) for donating the glycerol for this study.
Also, thanks to the staff at the Dairy Research Center on the University of Georgia (Tifton Campus)
for their assistance with animal care and sample collection and to Melissa Tawzer and Melanie
Renney (University of Georgia Animal and Dairy Science Department) for their laboratory and
technical assistance.

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PII: S0022-0302(12)00856-9

doi:10.3168/jds.2012-5760

© 2013 American Dairy Science Association. Published by Elsevier Inc. All rights reserved.

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Journal of Dairy Science

Volume 96, Issue 1 , Pages 470-476, January 2013